Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13478–13491

Small cat surveys: 10 years of data from Central Kalimantan, Indonesian Borneo

Karen Anne Jeffers¹, Adul ² & Susan Mary Cheyne³

^1-3Borneo Nature Foundation, Jalan Bukit Raya Induk No. 17, Palangka Raya, 73112, Central Kalimantan, Indonesia.

³Oxford Brookes University, Department of Social Sciences, Faculty of Humanities and Social Sciences, Gibbs Building, Headington Campus, Oxford, OX3 0BP, UK.

¹k.jeffers@borneonature.org, ²adul.outrop@gmail.com, ³s.cheyne@borneonature.org (corresponding author)

Abstract: We present an update on the photographic detections from camera traps and the activity patterns of Borneo’s four small cats, namely, Sunda Leopard Cat Prionailurus javanensis, Flat-headed Cat P. planiceps, Marbled Cat Pardofelis marmorata, and Bay Cat Catopuma badia, at two sites in Central Kalimantan, Indonesia.  Camera trap survey data of 10 years (2008–2018) from the first site in Sebangau provide details about the temporal partitioning of these small cats from each other but overlap with Sunda Clouded Leopard Neofelis diardi.  The activity of Flat-headed Cat was higher after midnight and that of Leopard Cat at night with no clear preference before or after midnight.  The Marbled Cat is predominantly diurnal, but the remaining three cats have flexible activity periods.  While limited data are available from Rungan, the second site, we confirmed the presence of all four small cat species found on Borneo, though we have insufficient data to comment on the Bay Cat.  The cat sightings, however, are intermittent and may reflect the unprotected status of this forest.  Leopard Cats appear relatively unaffected by habitat disturbance based on encounter rates on camera traps.  Conservationists, both NGOs and the government, must pay particular attention to specialists like Flat-headed Cats and Bay Cats when assessing habitat suitability for long-term cat conservation.

Keywords: Activity patterns, camera traps, Catopuma badia, diversity, felids, fire, peat-swamp forest, Pardofelis marmorata, Prionailurus javanensis, Prionailurus planiceps.

Bahasa Indonesia Abstract: Kami menyajikan pembaruan pada pendeteksian fotografi dari perangkap kamera dan pola aktivitas empat kucing kecil Borneo yaitu Kucing Kuwuk Prionailurus bengalensis, Kucing Batu Pardofelis marmorata, Kucing Tandang Prionailurus planiceps dan Kucing Merah Cat Catopuma badia di dua lokasi di Kalimantan Tengah, Indonesia. Data survei perangkap kamera 10 tahun (2008–2018) dari situs pertama di Sebangau memberikan rincian tentang partisi sementara kucing kecil ini dari satu sama lain tetapi tumpang tindih dengan Macan Dahan Neofelis diardi. Aktivitas kucing kepala datar lebih tinggi setelah tengah malam, dan Kucing Kuwuk di malam hari tanpa preferensi yang jelas sebelum atau setelah tengah malam. Kucing Batu didominasi diurnal, tetapi ketiga kucing memiliki periode aktivitas yang fleksibel. Sementara data terbatas tersedia dari situs kedua (Rungan), kami telah mengkonfirmasi keberadaan keempat spesies kucing kecil yang ditemukan di Borneo, meskipun kami tidak memiliki cukup data untuk mengomentari Kucing Merah. Namun, penampakan kucing berselang-seling dan mungkin mencerminkan status hutan yang tidak terlindung. Kucing Kuwuk kembali muncul relatif tidak terpengaruh oleh gangguan habitat berdasarkan pada tingkat pertemuan pada perangkap kamera. Konservasionis (LSM dan pemerintah) harus memberi perhatian khusus kepada spesialis misalnya Kucing Tandang dan Kucing Merah ketika menilai kecocokan habitat untuk konservasi kucing jangka panjang.

Introduction

In the absence of Tiger Panthera tigris, Borneo’s cats represent a fascinating guild, and all are in need of conservation attention.  Five species occur on Borneo: the Sunda Clouded Leopard Neofelis diardi (Endangered (EN) on the IUCN Red List; Hearn et al. 2016c), the Bay Cat Catopuma badia (EN and endemic to Borneo; Hearn et al. 2016a), the Flat-headed Cat Prionailurus planiceps (EN; Wilting et al. 2016a), the Marbled Cat Pardofelis marmorata (Near Threatened; Ross et al. 2016a), and the Sunda Leopard Cat Prionailurus javanensis (Least Concern; Ross et al. 2016).They are all protected under Indonesian law (P.20/MENLHK/SETJEN/KUM.1/6/2018).  As they are rare and elusive, it is difficult to study them in the wild.  Thus, there is limited knowledge about their ecology despite increased scientific interest.

Since 2008, Borneo Nature Foundation (BNF) and University of Palangka Raya, Centre for the International Management of Tropical Peatlands (UPR-CIMTROP), have been conducting long-term monitoring of the Sunda Clouded Leopard Neofelis diardi in Indonesia and, specifically, N. d. borneensis in Central Kalimantan.  During this time, we also opportunistically obtained images of three of the four small cats in the Sebangau catchment area.  Given the disturbed mosaic nature of Sebangau, we also hope to determine which areas are unsuitable for small cats, either due to habitat changes and/or due to human disturbance.  Live Leopard Cats are more common in Kalimantan markets for sale as pets than Sunda Clouded Leopards, and their skins are more often found in homes (Rabinowitz et al. 1987).  Far less is known about small cat movements, habitat preferences, seasonal movements, breeding patterns, and effects of anthropogenic disturbance across their range.  These data are particularly lacking from tropical peat-swamp forests.  We present here updated information on temporal activity pattern of four small cats from the Central Kalimantan region of Indonesia.

Study areas

The Sebangau catchment, Central Kalimantan, Indonesia (Fig. 1), is a peat-swamp forest (mixed-swamp forest sub-type) covering an area of ~5,600km².  This study took place in the 50km² research forest located in the northeast.  The area was logged under a concession system between 1991 and 1997 followed by illegal logging between 1997 and 2004.  The site is at an altitude of about 10m.  The area was significantly affected by the forest fires that impacted Indonesia in 2015.

The second research site was established in 2016 in the Rungan Forest, which covers about 1,440km² between the Kahayan and Rungan rivers.  The forest is a lowland forest mosaic comprising peat-swamp, ‘kerangas’ (heathland) and the dominant canopy trees are Palaquium sp. (nyatoh), Syzygium sp. (jambu) and Shorea (meranti), (Dipterocarpaceae family).  BNF and the Wildlife Conservation Research Unit (WildCRU) of the University of Oxford initiated the Sebangau Felid Project in May 2008, and BNF initiated the Rungan work in 2016.

Material and Methods

Since 2008, a total of 210 camera locations were surveyed across both forest areas with an average of 30 units in each forest area at any one time.  Between 2008 and 2012, cameras were set up in pairs in Sebangau and subsequently as single units.  In the Rungan site, cameras were all set up as single units.  Cameras were set in a stratified random survey design.  Cameras were placed 500m to 1,000m apart and were in each location for a minimum of six months; some cameras were in the same location since May 2008.  Locations were selected to cover a range of habitats and disturbances within the forests, avoiding streams and slopes wherever present.  Camera traps were placed along established human-made trails (more than four years old) and, where possible, watering areas, to maximise the success rate of photographic captures.  A combination of camera models were used, including Cuddeback Expert®, Cuddeback Capture IR® (Cuddeback Digital, Non-Typical Inc, WI, USA) Maginon, Crenova, and Bushnell.  Cameras were checked every 40 days when batteries were changed and SD cards exchanged.  Data were managed in a custom Microsoft Access database.  Active behaviour times were calculated using the kernel density method (‘href’ bandwidth for kernel smoothing; Ridout & Linkie 2009; Meredith & Ridout 2016) to account for average dawn and dusk times in the sites, which are situated almost on the equator (for more information on the Sebangau study site, see Cheyne & Macdonald 2011; Cheyne et al. 2016b).  Detection rate was estimated as number of detections/100 trap nights.  Weather data were collected daily at each research site and fire data was obtained from the Indonesian Agency of Meteorology, Climatology, and Geophysics.  A 30-min interval between photos of the same species was used to determine if photos of were an independent event at the same location and date.

Results

The number of camera trap (CT) stations at each site varied annually due to broken units (Table 1).

Detailed descriptions of CT locations are in Appendices 1 & 2, including descriptions of the microhabitats, number of trap nights, number of detections, and careful descriptions of the setup around the CT stations.  CTs were placed in different habitats across the two sites (Table 2).

The small cats were recorded by 56 of 83 camera traps in Sebangau (67%) and by 16 of 37 camera traps in Rungan (43%) (Table 3).

All small cats in Sebangau were photographed in all main habitat types in the interior forest, <20m from the forest edge and in disturbed areas.  All four small cats in Rungan were recorded in the interior forest; only the Flat-headed Cat was recorded near the lake.

Sebangau

Since the cameras were first placed in May 2008, we captured 157 independent images of Sunda Clouded Leopards (Image 1), but only 109 of Sunda Leopard Cats, 54 of Marbled Cats (Image 2), and 33 of Flat-headed Cats (Image 3).  Compared to the average detection rates of small cats since the inception of the camera trap study in 2008, there was a decline in the detections of Marbled Cats and Flat-headed Cats (Fig. 2).  From 2014 to July 2018, there was an average of 3.9 independent Marbled Cat images/month (min=0, max=16).  No Flat-headed Cats were recorded by camera traps between January 2014 and February 2018 (Fig. 2), which coincided with a significant fire event from September to November 2015.

Of the three small cat species, the Sunda Leopard Cat  is predominantly nocturnal with no clear preference for time of night.  Flat-headed Cats also showed nocturnal activity but with a slight preference for post-midnight hours.  Marbled Cats are strongly diurnal.  Interestingly, 65% of 115 nocturnal records (18.00–05.59 h) of Sunda Clouded Leopards were between 01.00h and 05.59h (Fig. 3), thus overlapping with the preferred active time for Flat-headed Cats.

Rungan

Three of the small cats were confirmed in Rungan in the first few months of the study but it took 12 months to confirm the presence of the Flat-headed Cat (Table 4).

Bay Cat

With over eight years of long-term camera trap surveying in the peat-swamp forest of the Sebangau catchment, the Bay Catwas not detected, and ongoing work suggests that it is not found in peat-swamp forests.  The Bay Cat was confirmed in mosaic heath/ peat-swamp forest habitat for the first time (Sastramidjaja et al. 2015; Cheyne et al. 2016a, 2017).  Through the use of camera traps, we present new location information on the distribution of Bay Cat in Kalimantan.  This new location is approximately 64km southeast outside the range depicted by Hearn et al. (2016a).  Our record of Borneo Bay Cat from the new habitat (heath/ peat-swamp forest) warrants further surveys in different habitat types to fully understand Bay Cat distribution and ecologic needs.

Table 1. Number of camera trap stations per year in the study site.

Table 2. Number of camera trap (CT) locations at each site (NA - habitat type not present in the study site).

Table 3. Camera trap (CT) locations with detections and non-detections of small cats with mean occupancy estimates (Ψ) in the study sites. NA indicates occupancy cannot be calculated due to no small cats being photographed at these locations.

Table 4.  Capture rate of small cat species from the inception of the camera trap surveys in Rungan between June 2016 and May 2018.­

Discussion

The small cats are appearing evenly across the habitat types in both Sebangan and Rungan, with the exception of the Bay Cat that likely does not exist in deep ombrogenous peat-swamp forest (Sebangau).  Additionally, we have evidence of breeding in Flat-headed Cat and Marbled Cat in Sebangau (images of kittens) (Images 4 & 5).

Flat-headed Cats have a more irregular capture rate and though they are active throughout the day, more captures are obtained at night and therefore they are predominantly nocturnal.  Leopard Cats have a more regular capture rate and appear to be active both during the day and night, though they appear to avoid the hottest time of the day (11.00–13.00 h).  Marbled Cats have a regular capture rate with the majority of images taken during the day (05.00–16.00 h), suggesting they are diurnal.  There is only one image of a Bay Cat taken at 11.17h.  These data are similar to those of Hearn et al. (2018), though these authors did not obtain sufficient images of Flat-headed Cats to make a detailed analysis.

Peat-swamp and associated lowland wetlands are postulated to be an important habitat for Flat-headed Cats (Cheyne et al. 2009; Wilting et al. 2010, 2016b; Cheyne & Macdonald 2011; Adul et al. 2015).  Marbled Cats are not believed to frequent roads or plantations (Hearn et al. 2016c) and prefer intact forests, though data are lacking on this cat (Rustam et al. 2016).

Peat-swamp and associated lowland wetlands were suggested to be poor or marginal habitat for Sunda Leopard Cat (Mohamed et al. 2016), but our work suggests that Sunda Leopard Cat are far more common (Cheyne & Macdonald 2011; Adul et al. 2015; Cheyne et al. 2016b).

The infrequent capture of the small cats in both sites is likely an artefact of the placing of the cameras (±1km apart) to focus on the wide-ranging Sunda Clouded Leopard.  By moving the cameras closer (±500m) we hope to determine the population density for the small cats, determine if the Bay Cat is indeed absent from this forest, and to continue our monitoring of the Sunda Clouded Leopard population.  The long period of time required to obtain images of small cats, possibly due to the placement of the cameras targeting Sunda Clouded Leopard, highlights the importance of long-term data and monitoring to avoid false-negative presence data.  Sunda Leopard Cat is the most commonly recorded species in the study site.  Marbled Cat is hard to study and, as many are arboreal, having cameras mainly on the forest floor means we could be missing out on key aspects of their behaviour.  Flat-headed Cat is a wetland specialist and prefers forests with water (Wilting et al. 2010).  Its diet likely consists of fish, frogs, and small mammals, and it may fill a niche on Borneo filled by the Fishing Cat Prionailurus viverrinus elsewhere in Asia (Iwaniuk et al. 2001).  Due to this dependence on wetlands, we think that the devastating fires of 2015 may have severely impacted the Flat-headed Cat population, pushing it away from fire-affected areas.  Our preliminary results suggest that Flat-headed Cats are returning to these areas, which BNF is actively working to restore.  These data could be an artefact of survey effort (a high number of broken cameras) or a genuine reflection of this species behaviour.

Large parts of the peat-swamp are naturally seasonally flooded for up to nine months per year.  The flooded nature of Sebangau does not always prevent the animals using the ground; indeed we have evidence of male Orangutans Pongo pygmaeus wading through water (Ancrenaz et al. 2014).  Keeping a selection of camera trap locations the same over several months or years allows for variations in detection to be accounted for; given the regular flooding of the forest, it is likely that the wildlife is accustomed to this.  We do notice animals using our boardwalks more regularly in the wet season.  Peatlands and associated forest fires are a crucial conservation concern in Kalimantan (Gaveau et al. 2016; Miettinen et al. 2017).  This is especially true during dry years such as in 2015 when a strong El Niño event led to particularly dry conditions.  From August to November of that year, MODIS satellites detected over 50,000 fire hotspots in Kalimantan, 53% of which were on peatland (Gaveau et al. 2016; Miettinen et al. 2017).  Since 2006, 17.35% of forest in the core Sebangau research area burned down (9.63% in 2015 alone—5.3km² of 55km²).  Of particular threat to the Flat-headed Cat are peat drainage and drying out due to logging canals, the loss of permanent water, and increased hydrologic instability (Page et al. 2009; Vanthomme et al. 2013).

Conservation impact

This work represents the first-ever comprehensive and long-term survey of small cats in Central Kalimantan.  There is a severe lack of data on these species in non-protected or small forest areas that may also contain viable populations.  It is crucial to remember that, while these surveys indicate the continued presence of these cats, habitat loss, wildlife trade, and likely presence of populations in non-protected areas means that more work is needed to understand the impacts of anthropogenic activities on these cats.  As detailed in Appendix 1, this project provides extensive and detailed data about many wildlife species in Sebangau and Rungan forests in addition to the cats—an additional 7,959 images (2,765 videos) of 74 species.  Of these, two are IUCN Red Listed as Critically Endangered, five as Endangered, 14 as Vulnerable, 12 as Near Threatened, and 41 as Least Concern.

For images / figures – click here

References

Adul, B. Ripoll, S.H. Limin & S.M. Cheyne (2015). Felids of Sebangau: camera trapping to estimate activity patterns and population abundance in Central Kalimantan, Indonesia. Biodiversitas 16(2): 151–155;  https://doi.org/10.13057/biodiv/d160208

Ancrenaz, M., R. Sollmann, E. Meijaard, A.J. Hearn, J. Ross, H. Samejima, B. Loken, S.M. Cheyne, D.J. Stark, P.C. Gardner, B. Goossens, A. Mohamed, T. Bohm, I. Matsuda, M. Nakabayasi, S.K. Lee, H. Bernard, J. Brodie, S. Wich, G. Fredriksson, G. Hanya, M.E. Harrison, T. Kanamori, P. Kretzschmar, D.W. Macdonald, P. Riger, S. Spehar, L.N. Ambu & A. Wilting (2014). Coming down from the trees: is terrestrial activity in Bornean orangutans natural or disturbance driven? Scientific Reports 4: 1–5.

Cheyne, S.M., Adul, F. Veen, J.F. Van, B.R. Capilla, N. Boyd & S. Maimunah (2017). First record of the Bay Cat in mosaic heath/peat-swamp forest, Kalimantan, Indonesia. Cat News 65: 48.

Cheyne, S.M. & D.W. Macdonald (2011). Wild felid diversity and activity patterns in Sabangau peat-swamp forest, Indonesian Borneo. Oryx 45(1): 119–124.

Cheyne, S.M., S. Maimunah & F. van Veen (2016a). Camera traps reveal wonders in Ironwood Forests. Peatlands International 3: 14–15.

Cheyne, S.M., H. Morrogh-Bernard & D.W. Macdonald (2009). First Flat-headed Cat from Sabangau peat-swamp forest, Indonesian Borneo. Cat News 51: 18.

Cheyne, S.M., W.J. Sastramidjaja, Muhalir, Y. Rayadin & D.W. Macdonald (2016b). Mammalian communities as indicators of disturbance across Indonesian Borneo. Global Ecology and Conservation 7: 157–173; https://doi.org/10.1016/j.gecco.2016.06.002

Gaveau, D.L.A., D. Sheil, Husnayaen, M.A. Salim, S. Arjasakusuma, M. Ancrenaz, P. Pacheco & E. Meijaard (2016). Rapid conversions and avoided deforestation: examining four decades of industrial plantation expansion in Borneo. Scientific Reports 6: 32017.

Hearn, A., J. Brodie, S. Cheyne, B. Loken, J. Ross & A. Wilting (2016a). Catopuma badia. In: The IUCN Red List of Threatened Species: e.T4037A112910221. Accessed on 14 October 2018. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T4037A50650716.en

Hearn, A.J., S.A. Cushman, J. Ross, B. Goossens, L.T.B. Hunter & D.W. Macdonald (2018). Spatio-temporal ecology of sympatric felids on Borneo. Evidence for resource partitioning? PLoS One 13(7): e0200828.

Hearn, A.J., J. Ross, H. Bernard, S.A. Bakar, L.T.B. Hunter & D.W. Macdonald (2016b). The first estimates of Marbled Cat Pardofelis marmorata population density from Bornean primary and selectively logged forest. PLoS One 11(3): e0151046.

Hearn, A., J. Ross, J. Brodie, S. Cheyne, I.A. Haidir, B. Loken, J. Mathai, A. Wilting & J. McCarthy (2016c). Neofelis diardi. In: The IUCN Red List of Threatened Species: e.T136603A97212874. Accessed on 14 October 2018. https://doi.org/10.2305/IUCN.UK.2015-4.RLTS.T136603A50664601.en

Iwaniuk, A.N., W.G. Blankstein & I.Q. Whishaw (2001). Observations of the feeding behaviour of Fishing Cats (Prionailurus viverrinus). Mammalia 65(1): 89–91.

Meredith, M. & M. Ridout (2016). Overview of the overlap package. In: R Development Core Team, R Package. 3.3.1. Edition. R Foundation for Statistical Computing, Vienna, Austria; https://cran.r-project.org/web/packages/overlap/vignettes/overlap.pdf

Miettinen, J., C. Shi & S. Liew (2017). Fire distribution in peninsular Malaysia, Sumatra and Borneo in 2015 with special emphasis on peatland fires. Environmental Management 60(4): 747–757.

Mohamed, A., J. Ross, A.J. Hearn, S.M. Cheyne, R. Alfred, H. Bernard, R. Boonratana, H. Samejima, M. Heydon, D.M. Augeri, J.F. Brodie, A. Giordano, G. Fredriksson, J. Hall, B. Loken, Y. Nakashima, J.D. Pilgrim, Rustam, G. Semiadi, T. van Berkel, J. Hon, N.T.L. Lim, A.J. Marshall, J. Mathai, D.W. Macdonald, C. Breitenmoser-Würsten, S. Kramer-Schadt & A. Wilting (2016). Predicted distribution of the Leopard Cat Prionailurus bengalensis (Mammalia: Carnivora: Felidae) on Borneo. Raffles Bulletin of Zoology, Supplement 33: 180–185.

Page, S., A. Hoscilo, H. Wosten, J. Jauhiainen, M. Silvius, J. Rieley, H. Ritzema, K. Tansey, L. Graham, H. Vasander & S. Limin (2009). Restoration ecology of lowland tropical peatlands in southeast Asia: current knowledge and future research directions. Ecosystems 12(6): 888–905.

Rabinowitz, A., P. Andau & P.P.K. Chai (1987). The Clouded Leopard in Malaysian Borneo. Oryx 21: 107–111.

Ridout, M. & M. Linkie (2009). Estimating overlap of daily activity patterns from camera trap data. Journal of Agricultural, Biological, and Environmental Statistics 14(3): 322–337.

Ross, J., J. Brodie, S. Cheyne, A. Datta, A. Hearn, B. Loken, A. Lynam, J. McCarthy, C. Phan, A. Rasphone, P. Singh & A. Wilting (2016a). Pardofelis marmorata. In: The IUCN Red List of Threatened Species: e.T16218A97164299. Accessed on 14 October 2018. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T16218A97164299.en

Ross, J., J. Brodie, S. Cheyne, A. Hearn, M. Izawa, B. Loken, A. Lynam, J. McCarthy, S. Mukherjee, C. Phan, A. Rasphone & A. Wilting (2016b). Prionailurus bengalensis. In: The IUCN Red List of Threatened Species: e.T18146A50661611. Accessed on 14 October 2018. https://doi.org/10.2305/IUCN.UK.2015-4.RLTS.T18146A50661611.en

Rustam, R., A.J. Hearn, J. Ross, R. Alfred, H. Samejima, M. Heydon, S.M. Cheyne, J.F. Brodie,  A.J. Giordano, H. Bernard, R. Boonratana, B. Loken, A. Mohamed, M.-A. Jayasilan,  D.M. Augeri, J. Eaton, J. Hon, A.J. Marshall, J. Mathai, G. Semiadi, D.W. Macdonald, C. Breitenmoser-Würsten, S. Kramer-Schadt & A. Wilting (2016). Predicted distribution of the Marbled Cat Pardofelis marmorata (Mammalia: Carnivora: Felidae) on Borneo. Raffles Bulletin of Zoology, Supplement 33: 157–164.

Sastramidjaja, W.J., S.M. Cheyne, B. Loken & D.W. Macdonald (2015). The Bay Cat in Kalimantan, new information from recent sightings. Cat News 62: 10–12.

Vanthomme, H., J. Kolowski, L. Korte & A. Alonso (2013). Distribution of a community of mammals in relation to roads and other human disturbances in Gabon, central Africa. Conservation Biology 27(2): 281–291.

Wilting, A., J. Brodie, S. Cheyne, A. Hearn, A. Lynam, J. Mathai, J. McCarthy, E. Meijaard, A. Mohamed, J. Ross, S. Sunarto & C. Traeholt (2016a). Prionailurus planiceps. In: The IUCN Red List of Threatened Species: e.T18148A50662095. Accessed on 14 October 2018. https://doi.org/10.2305/IUCN.UK.2015-2.RLTS.T18148A50662095.en

Wilting, A., S.M. Cheyne, A. Mohamed, A.J. Hearn, J. Ross, H. Samejima, B. Boonratana, A.J. Marshall, J.F. Brodie, A. Giordiano, J.A. Eaton, J. Hall, J.D. Pilgrim, M. Heydon, G. Semiadi, E. Meijaard, D.W. Macdonald, C. Breitenmoser-Würsten & S. Kramer-Schadt (2016b). Predicted distribution of the Flat-headed Cat Prionailurus planiceps (Mammalia: Carnivora: Felidae) on Borneo. Raffles Bulletin of Zoology, Supplement 33: 173–179.

Wilting, A., A. Cord, A.J. Hearn, D. Hesse, A. Mohamed, C. Traeholdt, S.M. Cheyne, S. Sunarto, M.-A. Jayasilan, J. Ross, A.C. Shapiro, S. Dech, C. Breitenmoser, J.W. Duckworth, J. Sanderson & J. Hofer (2010). Modelling the species distribution of Flat-Headed Cats (Prionailurus planiceps), an Endangered south-east Asian small felid. PLoS One 5(3): e9612.

Appendix 1. Summary of camera trap (CT) locations in Sebangau, Central Kalimantan Indonesian Borneo. LC - Leopard Cat, MC - Marbled Cat, FHC - Flat-headed Cat.